Zootaxa 287: 1–54 (2003) ISSN 1175-5326 (Print Edition) ZOOTAXA 287 Copyright © 2003 Magnolia Press ISSN 1175-5334 (Online Edition)

Seven new species of the Neotropical electric fish Gymnotus (Teleo- stei, Gymnotiformes) with a redescription of G. carapo (Linnaeus)

JAMES S. ALBERT* & WILLIAM G.R. CRAMPTON Florida Museum of Natural History, University of Florida, Gainesville, FL 32611 USA * Corresponding author; E-mail. [email protected]

Table of contents

Abstract ...... 1 Introduction...... 2 MaterialsandMethods ...... 4 Monophyly and Species-Groups of Gymnotus ...... 6 SystematicDescriptions...... 7 Gymnotus carapo Species-Group(Albert2001)...... 7 Gymnotus choco n.sp.Albert,Crampton,andMaldonado...... 26 Gymnotus esmeraldas n.sp.AlbertandCrampton ...... 30 Gymnotus henni n.sp.Albert,Crampton,andMaldonado ...... 32 Gymnotus paraguensis n.sp.AlbertandCrampton ...... 34 Gymnotus tigre n.sp.AlbertandCrampton ...... 36 Gymnotus pantherinus Species-Group(Albert2001) ...... 39 Gymnotus javari n.sp.Albert,CramptonandHagedorn...... 39 Gymnotus panamensis n.sp.AlbertandCrampton...... 43 Discussion ...... 46 Summary ...... 47 Acknowledgements...... 49 References...... 50 Appendix...... 53

Abstract

Seven new species of Gymnotus are described, and a redescription of the type species G. carapo sensu-stricto Linnaeus (G. carapo s.s.) is provided, from examination of populations from through- out tropical South and Middle America. The new species are described on the basis of unique com- binations of characters. Five of the new species are members of the G. c ar ap o species-group: 1, Gymnotus choco n. sp., from the Baudó and Atrato basins on the Pacific and Caribbean slopes of Colombia; 2, Gymnotus esmeraldas n. sp., from the Esmeraldas and Guayaquil basins on the Pacific Slope of Ecuador; 3, Gymnotus henni n. sp., from the Calima and Juradó basins on the Pacific Slope of Colombia; 4, Gymnotus paraguensis n. sp., from the Paraguay basin; 5, Gymnotus

Accepted: 22 August 2003; published: 5 September 2003 1 ZOOTAXA tigre n. sp., from the Amazon basin of Ecuador, Peru and Brazil. Two of the new species are mem- 287 bers of the G. pantherinus species-group: 1, Gymnotus javari n. sp., from the Amazon, Napo, Jav- arí, and Ucayali basins; 2, Gymnotus panamensis n. sp., from Atlantic slope of western Panama. Gymnotus carapo s.s. is restricted to the Amazon and Orinoco basins, the Island of Trinidad, the coastal basins of the Guianas, and the Brazilian state of Piauí. Six allopatric populations of G. c a r- apo s.s are recognized from differences in the mean, modal or median values of morphometric and meristic traits: 1, Eastern Amazon; 2, Parnaíba and Itapicuru basins; 3, Branco basin; 4; Guiana Shield, Orinoco basin and Island of Trinidad; 5, Madeira basin of Brazil, Bolivia and Peru; 6, West- ern Amazon basin of Brazil, Colombia, Ecuador, and Peru. Four of the new species representing both the G. c a ra po and G. pantherinus species-groups inhabit waters to the west of the Andean Cor- dillera. The biogeographic distributions of these species-groups suggest origins and early diversifi- cation before the most recent Andean orogeny, c. 8–12 mya.

Key words: biodiversity, classification, comparative morphology, species descriptions, South America

Introduction

The Neotropical electric fish genus Gymnotus (Linnaeus 1758) is substantially more diverse than was previously recognized. New methods of sampling, identifying, and col- lecting electric fishes have unveiled numerous species in previously unexplored habitats and regions (Crampton 1996a; Lundberg et al.1996; Crampton 1998; Albert & Crampton 2001). Further, the use of new techniques for characterizing morphology, electric organ discharges, and genetic differences have demonstrated that much of what was once regarded as intraspecific variation represents interspecific differences (Campos-da-Paz & Costa 1996; Fernandes-Matioli et al. 1998a; Fernandes-Matioli et al. 1998b; Albert et al. 1999; Fernandes-Matioli et al. 2000; Fernandes-Matioli et al. 2001; Albert & Crampton 2001; Fernandes-Matioli & Almeida-Toledo, 2001). Gymnotus species are aggressive nocturnal predators of fishes and other small aquatic animals, and most are territorial. The males of at least two Gymnotus species build and guard nests of foam and/or aquatic vegetation (Crampton & Hopkins pers. obs.). The type species Gymnotus carapo is reported to mouth brood its eggs and larvae (Kirschbaum & Wieczorek 2002). Gymnotus is the most geographically widespread of all gymnotiforms, extending from the Río Salado in the Pampas of Argentina (36°S) to the Río San Nicolás of southeastern Chiapas, Mexico (18°N), and is present in the continental waters of all South and Middle American countries except Chile and Belize (Albert 2001). Prior to this report, 25 species names were available for Gymnotus, of which five were synonymized with the type species G. carapo (Mago-Leccia 1994), and one with G. c o at es i (Campos-da- Paz 2000). The geographical ranges of the currently valid Gymnotus species are summa- rized in Table 1. Gymnotus exhibits its greatest diversity in the western portion of the Amazon basin where 10 species are currently recognized. Gymnotus anguillaris Hoede- man is recognized here as distinct from G. c o ro pi n ae Hoedeman with which it has been